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Home , Abyssobrotula, Abyssobrotula galatheae, Anoplogaster cornuta, Beryciformes, Cutthroat eel, Deep sea fish

, 5 ex-situ Ex-situ column and ly. tally. ater izon In-situ observations , David Shale 4 veillance, and (5) 7,1 sur tically and hor ution in the w (Contact e-mail: [email protected]) in-situ , Jørgen G. Nielsen , Jørgen 3 ces, both ver izontal distrib tan acking, (4) & Rupert Wienerroither & Rupert 6 Abstract Introduction Introduction perspective tical and hor , Pascal Lorance 2 acoustic tr cale intervals and dis Flat 12, Beachside Court, 14 Victoria Avenue, Swanage, Dorset, UK Avenue, Victoria Flat 12, Beachside Court, 14 5 ling, (3) identifies ver Instituto Canario de Ciencias Marinas, P.O.Box 56, 35200 Telde, Gran Canaria, Spain Telde, 56, 35200 Marinas, P.O.Box Instituto Canario de Ciencias Marsh J. Youngbluth Marsh J. 2 Harbor Branch Oceanographic Institution, 5600 US 1 North, Fort Pierce, FL 34946, USA Fort Pierce, FL Harbor Branch Oceanographic Institution, 5600 US 1 North, 6 , Fernando Bordes 1 ield samp Institute of Marine Research, P.O. Box 1870 Nordnes, N-5817 Bergen, Norway 1870 Nordnes, N-5817 Bergen, Box P.O. Institute of Marine Research, 1 mall to large-s e d'Yeu, B.P. 21105, 44311 Nantes Cedex 03, France 44311 21105, B.P. îe d'Yeu, IFREMER, Déartement Ecologie et Modèes pour l'Halieutique rue de l' 3 Department of Organismic Biology, and Biodiversity Research Unit, Hellbrunnerstrasse 34, A-5020 Salzburg, Austria A-5020 Salzburg, Research Unit, Hellbrunnerstrasse 34, Ecology and Biodiversity Biology, Department of Organismic 7 Zoological Museum, Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100, Copenhagen Ø, Denmark. Zoological Museum, Natural History Museum of Denmark, 4 Behavior and habitat selection of deep- : a methodological deep-sea fishes: selection of and habitat Behavior Franz Uiblein orphology, (2) f election over s depth, their systematics, ecology, and still requires intensified research. depth, their systematics, ecology, With With acoustic instruments vertical migrations and segregation patterns can be tracked. deployments of stationary gear document aggregation and diversity patterns on or close to the seafloor. with cameras and attraction devices or from underwater vehicles focus studies on behavior and habitat there and what are their behavioral habits? Although fishes are among the best studied fauna at greater experimentation. Functional morphology, often a “by-product” of taxonomic and systematic strongly studies, comparative is and correlative, and serves to examine hypotheses on behavioral sampling strategies. Field with towed gear complete understanding of behavioral diversity and to environmental factors. complete understanding of behavioral diversity and adaptations to environmental basic questions related to the evolution of deep-sea organisms are unresolved, e.g., how many live in unpressurized aquaria. An integrated approach which combines various methods would provide a more m behavioral studies of deep-sea fishes brought to the surface have been conducted to a limited extent, mainly of the most promising approaches to investigate the behavioral ecology are reviewed: (1) functional The , i.e. the vast area of the oceans below 200 m water depth, is largely unexplored and s The behavior and habitat selection of deep-sea fishes has been studied with various methods. Five

Franz Uiblein 05 approach is strongly comparative and correlative emphasizing the testing of hypotheses about behavioral about hypotheses of testing the emphasizing correlative and comparative strongly is The approach studies. systematic and taxonomic of “by-product” a as often arises morphology Functional (1) Functionalmorphology least at that estimated (1997) al. et Koslow described. been have species deep-sea 2700 About str adih 19) Ceotcie esn fcltts eeto o fo, rdtr, r oseiis n the in conspecifics or predators, food, of detection facilitates sensing Chemo-tactile 1997). Haedrich, M ques environmental features. species those e.g., dwellers, deep-sea “ancient” as regarded be can taxa Some exist. species 1000 another immediate surroundings or allows predators to track prey via currents (e.g., Wilson and Smith, 1984; Baird 1984; Smith, Wilson(e.g., and currents via prey track to predators allows or surroundings immediate exploratory methods can now be used to study deep-sea fish behavior. We provide here an overview of the of overview behavior.an fish Wehere deep-sea provide study to used be now can methods exploratory perception and communication modalities based on the comparative examination of chemo-tactile organs chemo-tactile of examination comparative the on based modalities communication and perception Many deep-sea fishes have large (calcium carbonate structures in the ) that may that ear) inner the in structures carbonate (calcium otoliths large have fishes deep-sea Many & Jumper, 1995). surveillance, and (5) and surveillance, in-situ n ed bd, is o bar or fins, body, head, on are fishes Other (Myctophidae). or (Ceratoidei) like organs light possess that in thisfield. (4) tracking, acoustic (3) sampling, field (2) morphology, functional (1) approaches: significant most five morpholog ae slcie on-ecpin ucin Pxo, 00. ifrne i ooih om mn closely among form in Differences 2000). (Paxton, function sound-perception selective a have epsa ihs Adise, 93. clgcly de-e fse hv be gopd no open-water, into grouped been have fishes deep-sea Ecologically, 1953). Andriashev, fishes; deep-sea related species strongly suggest that they permit intraspecific communication. For instance, the two sister two the instance, For communication. intraspecific permit they that suggest strongly species related pce o te the of species eai, n bto-soitd dmra seis Frhr et-eae sbiiin osdr as light also considers subdivision depth-related Further species. demersal bottom-associated, and pelagic, Atlantic at bathyal and abyssal depths (Uiblein et al., 2008). Recent morphological comparisons leading to leading comparisons morphological Recent 2008). al., et (Uiblein depths abyssal and bathyal at Atlantic bathypelagic m), -1000 (200 mesopelagic topography: bottom and maximally) m 1000 ca. (to penetration the resurrection of resurrection the m; 3500 ca. to (200 water,bathyal open the in species m) (>3500 abyssopelagic and m), 3500 ca. to (1000 et al., 2008). Members of this family have the capability to produce sounds based on “drumming” muscles “drumming” on based sounds produce to capability the have family this of Members 2008). al., et which from depth greatest bottom. The the on species m) 3500 (> abyssal and slope) lower upper,and mid- In recent years novel technologies have provided more frequent access and more detailed records in records detailed more and access frequent more provided have technologies novel years recent In a fishhasbeencollectedis8370m(Nielsen,1977). the the rde ad cusk and acrouridae, egies by by ategies h-eue ad yebrc r hyperbaric and light-reduced n aot ad about tions ucinl opooia suis f deep of studies morphological Functional y es differen less ically inves S. crassus S. s o xrm con extreme to aptations g h ad the tigating Spectrunculus , Oph , experimentation. As we hope, this review will stimulate more research more stimulate will review this hope, we experimentation. As ex-situ as a valid species have shown that they clearly differ in otolith shape (Uiblein shape otolith differin clearly they that shown have species valid a as s ad h str the and bels, ed and more more and tiated de se Marsh see idiidae; me of the deep sea, sea, deep the of egime e signif aptive and crassus S. (Ophidiidae), ons. An An ditions. ue f sound-produ of ucture sl r closely e f raim stru organism of icance ea fishes have commonly focused on on focused commonly have fishes -sea , 99 Cre ad uik 95 Me 1985; Musick and Carter 1979; all, d o s to elated n ary f s of array amazing ig t p it making oocr n h northern the in co-occur grandis, S. hallo g ras eg, n gr in (e.g., organs cing be o an to ossible w-w rs n r in ctures r s ater ing, ampling, s = s (= pecies n o dis to elation r fund swer analy n-visual non-visual ng, and and zing, ondary econdary ett and and rrett ers, enadiers, ntal amental nct tinct 06

Franz Uiblein , Neobythites family Ophidiidae, 39 (= 78 %) show coloration (Uiblein et al., 1994; Uiblein and Nielsen, 2005). This species shows typical occurs on the upper slope of the Northwestern Indian Ocean and the Red Sea and Neobythites Neobythites stefanovi hunting predators or deflecting their attacks to less vital parts of the body. Alternatively, they may Among 50 species of the cuskeel genus species is also provided (after Uiblein and Nielsen, 2005). 2005). species is also provided (after Uiblein and Nielsen, of species showing the respective color pattern and percentages referring to the total number of colored patterns. For each of the four different coloration types one species is shown as an example. The number The shallowest part of the deep sea is still weakly illuminated by downwelling light. Well-developed Neobythites Neobythites stefanovi Fig. 1. Fig. 1). Various functions have been ascribed to ocelli. They may reduce predation risk by, for instance, functions deterring for have instance, been ascribed They to may ocelli. reduce predation risk by, Various Fig. 1). circular circular to elliptical dark spots surrounded by a contrasting pale ring on the (Nielsen and Uiblein 1993; population, population, an upward-directed ontogenetic migration from about 800m to less than 600m was detected that ophidiid has been supported by investigations of eyespot (=ocellus) development and ontogenetic niche shifts in the has one ocellus on the dorsal fin positioned at about mid-body (Nielsen and Uiblein, 1993). In the Red Sea should be possible to ca. 500-600 m depending on water clarity (Clark and Denton 1961). The latter prediction should be able to perceive sunlight until a depth of around 1000 m and distinction of contrasting colors (Uiblein and Nielsen, 2005). Mann et al., 1997; Cruz and Lombarte, 2004; Rountree et al., 2006). Cruz and Lombarte, 2004; Rountree Mann et al., 1997; enhance enhance social communication and species recognition or they may serve several functions simultaneously differences in hearing capacity and selectivity may relate to the production of species-specific sound signals of species-specific to the production may relate capacity and selectivity in hearing differences and social communication in deep-sea fishes much like has been observed in shallow water species (e.g., visually that connect the swimbladder with modified ribs and the otic capsule (Merrett and Headrich, 1997). Species

Franz Uiblein 07 (2) Fieldsampling dev ets f a 10 m eutd n h cleto o mr ta 600 dl o jvnl fse blnig o 19 to belonging fishes juvenile or adult 68000 than more of collection the in resulted m 1000 ca. of depths conducted either with towed gear, such as various types of trawls, or with static gear such as longlines, baited longlines, as such gear static with or trawls, of types various as such gear, towed with either conducted omncto fr nacmn o seis eonto. iet vdne from evidence Direct recognition. species of enhancement for communication rp, r g or traps, bevtos s oee sil akn ad ute smln, seily n h Id-et aii, s edd to needed is improve theunderstandingofbothecologicalfunction ofthecolorationandsystematicsthisgenus. Pacific, Indo-West the in especially sampling, further and lacking still however is observations distribution and migration at larger scale, whereas static gear can be applied in studies of smaller scale habitat scale smaller of studies in applied be can gear static whereas scale, larger at migration and distribution indication for ocellus antipredator function, as risk from visually hunting predators should increase in shallower in increase should predators hunting visually from risk as function, antipredator ocellus for indication orders, 51families,126genera,and220species(Wienerroither 2005, Wienerroither etal.,2009). selection andaggregationformation. and moreilluminatedhabitats. diversity and distribution of pelagic fishes in the area of the Canary Islands (CE Atlantic). The Canary Islands Canary The Atlantic). (CE Islands Canary the of area the in fishes pelagic of distribution and diversity with the generally deeper-occurring population from the adjacent Red Sea revealed a significantly higher ocellus higher significantly a revealed Sea Red adjacent the from deeper-occurringpopulation generally the with n spatial enhance that slope steep a and shelf narrow a with origin volcanic of islands oceanic typical are imtr n h lte.Te nagd clu sol itniy h sgaig fet n ehne t vsblt to visibility its enhance and effect signaling the intensify should ocellus enlarged The latter. the in diameter neatos mn de-e ad daet hlo-ae hbtt (ili ad ods 19) ttl f 103 of total A 1999). Bordes, and (Uiblein habitats shallow-water adjacent and deep-sea among interactions pelagic trawl tows during seven cruises from above the island shelf and the slope and from the surface down to down surface the from and slope the and shelf island the above from cruises seven during tows trawl pelagic predators at larger, less illuminated depths (Uiblein, 1995). While this observation fits well with the assumed the with well fits observation this While 1995). (Uiblein, depths illuminated less larger, at predators ipu dumer Diaphus aggregations close to the shelf at night. High densities occurred particularly in areas of high and productivity high of areas in particularly occurred densities High night. at shelf the to close aggregations (W antipredator function of the ocellus, a posterior shift in the position of this color structure found in the Gulf of Gulf the in found structure color this of position the in shift posterior a ocellus, the of function antipredator Aden population required alternative interpretations. In a subsequent study of coloration patterns 50 patterns coloration of study subsequent a In interpretations. alternative required population Aden species, 22 species were found to have well-developed ocelli (Uiblein and Nielsen, 2005, Fig. 1). Among 1). Fig. 2005, Nielsen, and (Uiblein ocelli well-developed have to found were species 22 species, oocrig pce cnieal vrain n clu psto ad ubr curd hc sget a social a suggests which occurred number and position ocellus in variation considerable species co-occurring omncto fnto ta wud ii itrpcfc neatos n hbiiain Uben n Nielsen, and (Uiblein hybridization and interactions interspecific limit would that function communication 2005). on- ienerr ment of the the of elopment rc mu metric Field sampling of deep-sea fishes can also reveal important information on behavior. Sampling can be can Sampling behavior. on information important reveal also can fishes deep-sea of sampling Field y mul By apig ih oe ga hs epd o eie irto pten a eepiid y tde o the of studies by exemplified as patterns migration define to helped has gear towed with Sampling A morphological comparison of the of comparison Amorphological Thus, ocelli in ocelli Thus, r 05 W 2005, oither illn en te niao seis o vrial mgaig eoeai fse and fishes mesopelagic migrating vertically for species indicator the being gaussi Lepidophanes . Samp ets. ltid rae lsiiain n or and classification tivariate ilii ninl s imensional o te ru o hor of group the for lu (Uib ocellus Neobythites iene ng with towed towed with ling ihr t l, sub al., et rroither n) or ecolog four caling) n t l, 94 U 1994; al., et lein may serve two different functions, minimization of predation risk and social and risk predation of minimization functions, different two serve may ic pelagic ntal izontal specimens collected in the Gulf of Aden, Indian Ocean, Indian of Aden, Gulf the in collected specimens stefanovi N. dina ed). Other Other mitted). r gener gear n met tion migr ly separable separable ically i, 19 iblein, ors ators ds (hierarchical (hierarchical hods ly ally tween between lante ). This observation was interpreted as an an as interpreted was observation This 96). lows allows fishes rnfishes oups could be distinguished with the the with distinguished be could groups c w oceanic inves ke like g ver tigating glomerative clustering and and clustering agglomerative or in-situ yohm hygomii Hygophum r ad h is the and aters l r horizo or tical behavioral ex-situ Neobythites d s land

rmed formed lf helf al ntal

08

Franz Uiblein occurred Scomber colias collected during two cruises in the (Moridae), Mora with moro advanced gonadal Hygophum hygomii in the western North Pacific were tracked by towed bongo-net Diaphus theta to 73 species and 38 families (Uiblein et al., 1996, 1998; Lorance et al., Among 2002). different Number of individuals of the lanternfish Stationary gear, such as longlines, baited traps, or gillnets, provide insights about small-scale habitat Stationary gear, Migrations of deep-sea fishes may also serve . For instance, long-distance horizontal Canary Islands, Eastern Central Atlantic (after Wienerroither, 2003). Wienerroither, Atlantic (after Canary Islands, Eastern Central maturity was collected off eastern Fuerteventura, one of the two islands studied, at around 1000 m depth Oyashio front. selection and aggregation formation. For example, during long-line cruises in the area of Lanzarote and Furteventura, Canary Islands, operations at depths between 18 and 1208 m collected a total of 1885 fishes spawning migration from the Oyashio Subarctic and Western waters into the transition waters crossing the November 1997, an unusually high number of deep-sea , the Canary Islands and associated hydrological variability. During two periods, in October 1995 and fronts. Larvae and juveniles were distributed in transition waters, whereas larger individuals were collected abundance of single species were found. These results reflect to some extent the particular topography of in the Oyashio and the Western Subarctic waters. These results indicate that this species performs a peak abundance of larvae was observed in July in the transition waters between the Oyashio and Kuroshio depth zones clear variations in fish total weight and species number, and richness the and relative diversity, fishes. Such horizontal migrations can be seen in the context of both foraging and predation-risk avoidance. fishes. Such horizontal migrations can movements of the lanternfish and midwater collections of larvae, juveniles, and adults during different seasons (Moku et al., 2003). The belonging that were observed to migrate from the shelf into the oceanic water in opposed direction to the mesopelagic specific specific hydrological conditions, like eddies and (Fig. 2; 2003). Among Wienerroither, the group of typically shelf-associated fishes also some epipelagic species like chub Fig. 2.

Franz Uiblein 09 of organisms related to this layer. In parallel, pelagic trawl samples from both the and layer scattering deep the both from samples trawl pelagic layer.parallel, this In to related organisms of (3) Acoustic tracking this of individuals few a only seasons, other during and depths lesser At 1998). 1996, al., et (Uiblein tagging has been successfully adopted to track migrations of demersal fishes at great depth (e.g., Bagley et Bagley (e.g., depth great at fishes demersal of migrations track to adopted successfully been has tagging ver fish deep-sea sp l, 94 Pid ad aly 20) Aosis a ehne te understanding the enhanced has Acoustics 2000). Bagley, and Priede 1994; al., seasonally restricted spawning aggregation. Males arrived earlier than females at the spawning grounds and grounds spawning the at females than earlier arrived Males aggregation. spawning restricted seasonally patterns overlarge depthrangesanddeepscatteringlayer formation(e.g.,Opdaletal.,2008). there appeared to be distinct microhabitat preferences related to a specific depth and sites close to a canyon a to close sites and depth specific a to related preferences microhabitat distinct be to appeared there deep scattering layer formed at ca. 400 to 700 m depths off various islands and the diurnal vertical migration with localupwellingactivity(Fig.3). Fig. 3. Fig. s ee agt n h sm ae. hs dt srnl sget h for the suggest strongly data These area. same the in caught were ecies Number of deep-sea cod, deep-sea of Number Acoustic Acoustic During the pelagic cruises in the Canary Islands (see also above) acoustics was employed to study the m depth, SE Fuerteventura, Canary Islands, Eastern Central Atlantic. Note the high numbers close numbers high the Note Central Atlantic. Eastern Islands, Canary Fuerteventura, SE depth, m to theentranceofacanyon-likesteeptopographicstructure (afterUibleinetal.,1998). ods methods hrzna migr tical/horizontal

uding echosounders, sonar, and and sonar, echosounders, including Mora moro,Mora ions, ations, collected at bottom longline stations at between 800 and 1200 and 800 between at stations longline bottom at collected l distr spatial ibu , n ag and tion, tic tagging tagging acoustic n for gregation o o a sp a of mation o investig allow f etcl migration vertical of . A mation. ly and and atially ion of of ation tic coustic 10

Franz Uiblein April th species, Lampanyctus nhancing the spatial exchange and opes thus e cattering layer that after sunrise moves downwards and, allow s L. pusillus were collected by both trawl tows, and hence their vertical rrow platforms and steep sl and itially formed sh scattering the layer fishes and therein reside close to the surface at night, as shown in recent studies Echogram taken off SE Fuerteventura, Canary Islands, during late night and morning of 12 Acoustics can also assist to track horizontal migrations of deep-sea fauna, especially when the 2003, Bordes et al., 2009). Lampanyctus) inserted that were collected by both tows (after Bordes et al., 1999; Wienerroither 1997, with vertical position of pelagic trawl tows and drawings of three myctophid species (genus shallow migration activity in this area and during this particular time period is defined. migration activity in this area and during this particular time period is defined. L. alatus, L. photonotus, subsequently, later subsequently, unites with the deep scattering layer. Lanternfishes like the three interaction between the neritic and oceanic fauna. The forces inciting such horizontal migrations are still clearly shows the in description of acoustic methods used see Bordes et al., 1999). The echogram taken during this period overlapping with the continuously formed deep scattering layer after April sunrise 1997 on (Fig. 12 4; for a of volcanic origin with na For instance, off Fuerteventura, trawl hauls were performed close to the surface late at night and at a depth fishes which remained at the surface during the night. Like the Canary Islands, the Hawaiian archipelago is the surface were taken to track and identify the species undergoing those migrations (Bordes et al., 1999). Fig. 4. the Canaries. The Hawaiian study indicated a horizontal mass migration onto the shelf by mesopelagic off off Oahu, Hawaii (Benoit-Bird Au, and 2006). In this case a towed camera was used to identify the faunal components, but this method did not provide taxonomic resolution like the pelagic trawl-supported study in

Franz Uiblein 11 (4) In-situsurveillance pred but unknown, topography around oceanic islands and shallow may enhance trapping of oceanic oceanic of trapping enhance may seamounts shallow and islands oceanic around topography ht lo drc observ direct allow that like which may be used as a locally aggregating food resource by shelf-dwelling and mesopelagic and shelf-dwelling by resource food aggregating locally a as used be may which copepods like olwn idvdas uig owr lcmto. hs ehd a b ue, o isac, o investigate to instance, for used, be can method This locomotion. forward during individuals following su fishes (Bordesetal.,1999;seealsochapter2). i. 5. Fig. pcfc oeet atrs Jnsn t l, 96 Moe 20) cags n oy ooain eg, from (e.g., coloration body in changes 2002), Moore, 1986; al., et (Janssen patterns movement specific method items sp over a larger area during vertical or horizontal transects (e.g., Yoklavich et al., 1999; Lorance et al., 2002; al., et Lorance 1999; al., Yoklavich(e.g., et transects horizontal or vertical during area larger a over Drazen et al., 2003; Uiblein et al., 2002, 2003, 2006; Lorance et al., 2006; Stein et al., 2006; Uiblein, 2009; Uiblein, 2006; al., et Stein 2006; al., et Lorance 2006; 2003, 2002, al., et Uiblein 2003; al., et Drazen Fig. 5). Prolonged5). Fig. observ re fo s from bmerged ecies (Is compos In-situ canyon, Gulf of Maine, NW Atlantic, in September 2002 (dives nr. 4471, 4475, 4479) and September and 4479) 4475, 4471, nr. (dives 2002 September in Atlantic, NW Maine, of Gulf canyon, 2004 2004 lig background) (inpartafterUiblein etal.,2006). Mobile platforms such as underwater vehicles allow tracking fishes and investigating their behavior their investigating and fishes tracking allow vehicles underwater as such platforms Mobile etcl itiuin f he fs tx osre drn mne sbesbe ie i Oceanographer in dives submersible manned during observed taxa fish three of distribution Vertical ows observ allows aacs saoe ver ht-shadowed d Schwarand v 47, 75 48) Te dark The 4788). 4785, 4779, (dive s ition es oftudies s and hips and dens , od o co or food, ation, ion of ation deep-seabehaviorfish tzlose, ationofsingle individ o o rec or ation l ae design layer tical w equipped ity ts of foraging b foraging of elements , Guennegan1975, Rannou,and1979, Wil (Pr iede, 1994; Co ig f eair Sainr plat Stationary behavior. of ording ith on may be be may mpetition rs n b and cameras es dives during during dives ates uals be performedbe can llins n becan aoe horizo -shadowed avior such as search, d search, as such ehavior et al., 1999; Henriques et ach to ait g h ms influ most the among ievedbystopping thev night- attr om both s bothfrom ad eod s record and act al ntal e s oprd o day to compared as time n & S & son er layer ms include include forms tationarymo and on, or etection, al., 2 s h bo the indicates th, 1984)esmith,and e ats ehicle 002; King l f ential nig ihs This fishes. cavenging ng of food of handling r ta are that landers ors. Abrupt Abrupt actors. electeds e p bile etal.,

me (white (white time m, the the ttom, latforms timating 2007). itesor 12

Franz Uiblein -1 enadier, cutthroat ) among sites were thern -1 oundnose gr r kaupi, - Body lengths moved s - Body lengths moved Category -1 nchus (Fig. 6). In nor ticus in the Bay of Biscay, NE Atlantic. NE of Biscay, in the Bay phobra in the Bay of Atlantic Biscay, NE (Uiblein et and body lengths moved s -1 Syna Nautile Entire body whitish - Head whitish, body with Entire body whitish - Head whitish, body red colour patches - Body and head in “typical” red color Inactive - Active - Arriving disturbed Arriving - Active Inactive - holding - Drifting - No locomotion - Station Forward movement - Slightly above bottom - Attached to bottom (> 1 total length) - High above bottom Well total lengths) above bottom (> 3 At far - At short distance No disturbance - distance Body undulations s Nautile eel, tethus atlan dives in the Bay of Biscay, NE Atlantic . NE dives in the Bay of Biscay, Hoplos cutthroat body undulations s tance, in atlanticus, see Lorance et al., 2002 1), and Table or the response to and , al., 2003) during submersible Images of selected fish species videotaped for behavioral studies (Uiblein et Fig. 6. Behavior/Characteristics (taxa) Behavior/Characteristics oides rupestris, Overview Overview of the categories of behavior and other characteristics of deep-sea fishes based on video recordings during dives with the manned submersible the manned submersible during dives with recordings In-situ observations with the manned submersible Disturbance response (all) Disturbance response kaupi ) Swimming speed ( Coloration (Hoplostethus atlanticus) Activity level (all) (all) Locomotion behavior (all) Position in differences differences in swimming speed (measured as Coryphaen have been also found as, for ins position above bottom, and locomotion mode (e.g., forward movement, station holding, and drifting, see Table 1) position above bottom, and locomotion mode (e.g., forward movement, station Table holding, and drifting, see small spatial scale, and interspecific variation. Differences between adjacent habitats in activity level, vertical al., 2003) have revealed novel insights into the behavioral ecology of deep-sea fishes, their natural habitat use at bait (Widder et al., 2005). et al., 2005). bait (Widder white to red in orange roughy, Table Table 1.

Franz Uiblein 13 lanternfishes) or bottom-associated fishes (e.g., snailfishes, Liparidae, or , Zoarcidae) that extend from extend that Zoarcidae) eelpouts, or Liparidae, snailfishes, (e.g., fishes bottom-associated or lanternfishes) envi (5) Ex-situexperimentation in plasticity behavioral of degree high a suggest strongly results These 2002). al., et (Uiblein too discovered, the shelfintodeepsea(e.g.,Robison,1973;McCoskerand Anderson, 1976;SakuraiandKido,1992). can be kept alive for longer periods (e.g., McCosker and Anderson, 1976; Miwa et al., this volume). Reports of Reports volume). this al., et Miwa 1976; and Anderson, McCosker (e.g., periods longer for alive kept be can omncto, r oil eair eg, ili, 92 mrm t l, 04 Wn e a. 20) Fr this For 2005). al., et Wong 2004; al., et Amorim 1992; Uiblein, (e.g., behavior social or communication, epsa ihs loig dutet o ml-cl sail r eprl lcutos n yrgah ad food and hydrography in fluctuations temporal or spatial small-scale to adjustment allowing fishes deep-sea curling-up behavior (Fig. 7) that has been observed in nature. This posture may be interpreted as an antipredator an as interpreted be may posture This nature. in observed been has that 7) (Fig. behavior curling-up availability. needs to be carefully investigated in order to document whether they alter naturally occurring behavior (Uiblein behavior occurring naturally alter they whether document to order in investigated carefully be to needs eald eairl bevtos n scesu eprmnain r hwvr cre Otn clmtzn was acclimatizing Often scarce. however are experimentation successful and observations behavioral detailed et al., 2002; 2003). Interestingly, the detailed analysis of disturbance behavior itself can also provide information provide also can itself behavior disturbance Interestingly,of 2003). analysis 2002; detailed al., the et strategy, as it could serve to mimic the form of a medusa (Robison, 2004) or make detection and / or attacking or / and detection make or 2004) (Robison, medusa a of form the mimic to serve could it strategy,as purpose one may study deep-sea fishes either in captivity in the ocean or “ or ocean the in captivity in either fishes deep-sea study may one purpose not sufficiently accomplishedandvirtuallynooronlyfew behavioralelementscouldbestudied. /synopsis.html; Uiblein et al., 2006) several fish species were collected with the manned submersible manned the with collected were species fish several 2006) al., et Uiblein /synopsis.html; n ecpul n rato cpblte o de-e fse a wl a o nnhmn nue environmental induced non- on as well as fishes deep-sea of capabilities reaction and perceptual on more difficult for predators. Alternatively, it might be also a biomechanical phenomenon allowing the fish to fish the allowing phenomenon biomechanical a also be might it Alternatively, predators. for difficult more vessel or on land. Both normal and pressurized tanks have been used to maintain deep-sea fishes. Unpressurized fishes. deep-sea maintain to used been have tanks pressurized and normal Both land. on or vessel influences onbehavior(Uiblein&Lorance,2007;Uiblein, 2009). Sealink float moreeasily, ormaysimplyreflect sleepingorresting anddrifting. tanks are useful for species that occur in shallow water such as vertically migrating mesopelagic fishes (e.g., fishes mesopelagic migrating vertically as such water shallow in occur that species for useful are tanks (McCosker and Anderson, 1976) and was one of the first deep-sea fish species in which feeding behavior was behavior feeding which in species fish deep-sea first the of one was and 1976) Anderson, and (McCosker n te the and beanii, Serrivomer October 2005 in Oceanographer canyon at 2293 ft, kept in a small glass tank on the RVthe on tank glass small a in kept ft, 2293 at canyon Oceanographer in 2005 October transferred to the Woods Hole Aquarium a few days later where it was kept alive and survived for three months three for survived and alive kept was it where later days few Woodsa the Hole Aquarium to transferred (until 16 January 2006). However, the specimen was not observed feeding over the entire period (John Galbraith, (John period entire the However,over 2006). feeding January observed 16 not (until was specimen the personal communication) and most probably died of starvation. Another short-lived specimen collected during collected specimen short-lived Another starvation. of died probably most and communication) personal the same cruise was kept just several days on board in a kreisel (Fig. 7). In this container the fish showed typical showed fish the container this In 7). (Fig. kreisel a in board on days several just kept was cruise the same mna ad or and ronmental n kp aie n ak, mn te te hatchetfish the them among tanks, in alive kept and xeiet udr otold odtos r ncsay n re t rlt te ifrnil nlecs of influences differential the relate to order in necessary are conditions controlled under Experiments There is growing evidence from scientific laboratories as well as from public aquaria that deep-sea fishes deep-sea that aquaria public from as well as laboratories scientific from evidence growing is There All o eape drn rcn cuss f te uf f an (.. http://at-sea.org/missions/maineevent6 (e.g., Maine of Gulf the off cruises recent during example, For Another species of the genus the of species Another methods, even deployed cameras without bait (Koslow et al., 1995), create disturbance that disturbance create 1995), al., et (Koslow bait without cameras deployed even methods, in-situ ganism-r d f elated n seie o te atr a cletd n 5 on collected was latter the of specimen One atlanticum. Melanostigma Melanostigma, M. pammelas, M. Melanostigma, r o fs for fish on actors g pr aging, ryoeeu aculeatus, Argyropelecus was kept in captivity for more than a year a than more for captivity in kept was or edator ” after collection on board of a of board on collection after ex-situ” e, avoidance, Seward Johnson I Johnson Seward h swote eel sawtoothed the lear g, ning, cogn Johnson n, ition, and 14

Franz Uiblein Careproctus Careproctus have also the Anoplogaster imental chambers Pressurized exper show much of their behavioral repertoire. (Liparidae) (Liparidae) (Sakurai and Kido, 1992), and in the robust bathypelagic , There have been also successful trials to keep fishes in pressurized tanks after previous decompression or Examples Examples of experimental approaches that may be applicable to studies of deep-sea fish behavior come transfer (e.g., Smiley and Drawbridge, 2007, Miwa et al., this volume). Many of these systems are still period with stepwise transfer to decreasing pressure may allow acclimatizing of deep-sea fishes to surface disadvantage disadvantage of restricted access for manipulations of environmental factors. Decompression over a longer already been emphasized earlier (Poulsen, 1971; Uiblein, 1996a, b; Uiblein, 2000). already been emphasized earlier (Poulsen, 1971; Uiblein, 1996a, b; Uiblein, 2000). for fish to biology has also some relevance for considering ecological and evolutionary processes in the deep-sea has under development (e.g., Damasceno-Oliveira et al., 2004) and often the space provided in the tanks is too small (Uiblein et al., 1992) and choosing between illuminated and dark habitats (Uiblein et al., 1996). That cave in-situ dwellers which provide insights into evolutionary steps undergo when adapting to lightless conditions fishes. Particularly interesting are comparisons between so-called secondary and the more adapted primary cave studies of cave behavior and habitat selection are very much advanced compared to those of deep-sea search and prey detection at distance or type, size and behavioral activity of the prey were not investigated. at distance or type, size and behavioral search and prey detection to elicit chemically and/or mechanically mediated feeding responses. Many important parameters like predator cuskeels (order ) live in both habitats (e.g., Parzefall, 1996, Uiblein, 1996a, b). Experimental transfer from the ship on land. Using dim red light for illumination the fish were stimulated at various body parts 1996, 2000). Caves provide light-reduced conditions similar to the deep sea, and some groups of fishes like the (Childress and Meek, 1973). In the latter study the fangtooth were kept in large containers in a cold room after from work on cave animals (e.g., Uiblein et al., 1992; Uiblein and Parzefall, 1993; Uiblein 1996a, b, c; Parzefall (Anoplogasteridae), (Anoplogasteridae), which was kept successfully in tanks for several days (Robison, 1973) and fed with shrimps rastrinus conditions (St. John, 2003). observed (Belman and Anderson, 1979). Feeding behavior was also observed in the snailfish,

Franz Uiblein 15 have been used already in combination, e.g., acoustics and sampling (Fig. 4), acoustics and acoustics 4), (Fig. sampling and acoustics e.g., combination, in already used been have (Benoit-Bird et al., 2003), or 2003), al., et (Benoit-Bird hge itgain f hs vros ehd t gi a oe opee itr o bhvoa dvriy and diversity behavioral of picture complete more a gain to methods various these of integration higher a to environmental conditions at large depth, an important prerequisite for further progress in deep-sea in progress further for prerequisite important an depth, large at conditions environmental to adaptation fish behavioralecology. as, for instance, eelpouts of the genus the of eelpouts instance, for as, o epsa otm (.. Mrl ad enr 17; ivreg t l, 97 Slebr ad os, 1994; Bossé, and Silverberg 1987; al., et Silverberg 1979; Wenner, and Markle (e.g., bottoms deep-sea to Robison, 2004; Uiblein et al., 2006; Fig. 5), can be successfully collected in these different habitats, and studied and habitats, different these in collected successfully be can 5), Fig. 2006; al., et Uiblein 2004; Robison, both in-situandex-situ use. ay olcin o de-e fse ae tl aatn dtie mrhlgcl tde ta promise that studies morphological exciting andoftencompletely newinsightsintolifeinthe deepsea. detailed awaiting still are fishes deep-sea of collections Many pursued. i. 7: Fig. Various methods are currently available for the study of deep-sea fish behavior. Some of these methods these of Some behavior. fish deep-sea of study the for available currently are Variousmethods Johnson Johnson species with the Johnson Sealink manned submersible and transfer to on-board keeping systems. The systems. keeping on-board to transfer and submersible manned Sealink Johnson the with species Melanostigma atlanticum, Melanostigma eelpout, Atlantic f uln-padditn bhvo ta hv be as osre i ntr, evn ms poal to probably most serving nature, in observed also been have that behavior curling-up-and-drifting of Some fish species may prove especially useful for behavioral studies within an integrative approach such approach integrative an within studies behavioral for useful especially prove may species fish Some reduce predation risk (photographs of eelpout by David Shale). This figure also shows images of two of images shows also figure This Shale). David by eelpout of (photographs risk predation reduce other species that were kept in small glass tanks, the hatchetfish the tanks, glass small in kept were that species other atohd eel sawtoothed vertical bodyorientationthatcanbealsoencountered s at f n nertv apoc, ai txnmc n sseai wr nes o e oe actively more be to needs work systematic and taxonomic basic approach, integrative an of part As lmss f epsa ih eair ee ahrd uig eet rie wt te V Seward RV the with cruises recent during gathered were behavior fish deep-sea of glimpses Ex-situ ad I (http II and I . Te atr eand n ha-on oiin vr eea hus a hours, several over position head-down a in remained latter The beanii. Serrivomer and in-situ ://at-s ea.or which occur widely distributed from near the ocean surface ocean the near from distributed widely occur which Melanostigma observation (Figs. 5,7). In the future it will be important to reach to important be will it future the In 5,7). (Figs. observation ex-situ Conclusions andoutlook g/missions was maintained in a Kreisel tank where it displayed elements displayed it where tank Kreisel a in maintained was /maineevent6/s in situ. ynopsis Argyropelecusaculeatus l), after after .html), ion of several several of collection observation in-situ and the Bean's the and 16

Franz Uiblein

tical atinus tance in a clic ver Anoplogaster ations on the 1999. Epiand onus man impor aguchi. their Barathr lation of cy deep-sea fish ater fishes and enabling the simu deep-w Priede. 1999. behavioural observ

phism in the econdary ater organisms dimor References (Pisces, Sternophychidae). Deep-Sea Res I 42:675-696. (Pisces, Sternophychidae). Deep-Sea Res exual ater and s deep-w Sternoptyx diaphana ncient Musick. 1985. S tudy of marine or freshw J.A. long-term s rrett, M. Miya, T. Moreno, F. Perez, A. Ramos, T. Sutton and M. Yam scavenging fauna of the Patagonian slope. J. Mar. Biol. Ass. U.K. 79: 965-970. Biol. scavenging fauna of the Patagonian slope. J. Mar. 456-468. : ). Copeia 1973 (3): 602-603. cornuta: Beryciformes). Copeia 1973 (3): 602-603. (Ophidiidae). Copeia 1985 (1): 69-75. Ciencias Marinas (ICCM), Gobierno de Canarias, Viceconsejeria de Pesca, 326 pp. Viceconsejeria Ciencias Marinas (ICCM), Gobierno de Canarias, Me micronekton community. Mar. Eco. Prog. Ser. 319. 1-14. Eco. Prog. Ser. Mar. micronekton community. measured in situ from a manned submersible. Deep-Sea Research I 50. 221-229. Deep-Sea measured in situ from a manned submersible. islas Canarias, durante las campañas realizadas a bordo del B/E "La Bocaina". Instituto Canario de Zoarcidae). Copeia. 1979 (2): 366-369. Zoarcidae). Copeia. 1979 (2): 366-369. location problem in Catálogo Catálogo de especies meso y batipelágicas. Peces, moluscos y crustáceos. Colectadas con arrastre en las for Seabight, North-East Atlantic Ocean. J. Mar. Biol. Ass. U.K. 74: 473-480. Ass. U.K. 74: 473-480. Biol. Ocean. J. Mar. Atlantic Seabight, North-East mesopelagic mesopelagic fishes, acoustic data, and SST images collected off Lanzarote, Fuerteventura, 5: 1-45. Téc. ICCM 04-97". Inf. "La Bocaina Canaria, Canary Islands, during cruise and Gran Fish Biol. 65: 1512-1525. males of three closely related Lake Malawi cichlid species. J. Fish Biol. 65: 1358-1371. J. Fish Biol. 65: species. Lake Malawi cichlid closely related males of three DC). zoogeographical zoogeographical analysis. In: Notes on special problems in . Moscow, Nauk Leningrad: USSR, Akad. Ikhiol. Kom. (translation from Ichthyol. Lab., Bureau of Comm. , Washington llins, M.A., C. Yau, C.P. Nolan, P.M. Bagley and I.G. Cruz, A. and A. Lombarte. 2004. Otolith size and its relationship with colour patterns and sound production. J. Co Clarke, G.L. and E.J. Denton. 1961. Light and animal life. In: Hill, M.N. (ed.), The Sea. New pp Wiley, York, Childress, J.J. and Meek. R.P. 1973. Observations on the feeding behavior of a mesopelagic fish ( Carter, H.J. and Bordes, F., F. Uiblein, R. Castillo, A. Barrera, J.J. Castro, J. Coca, J. Gomez, K. Hansen, V. Hernandez, N. Benoit-Bird, Benoit-Bird, K.J. and Au. W.W.L. 2006. Extreme diel horizontal migrations by a tropical nearshore resident Benoit-Bird, Benoit-Bird, K.J., Au, W.W.L. C.D. Kelley and C. Taylor. 2003. Acoustic backscattering by deepwater fish Belman, B.W. and M.E. Anderson. 1979. Aquarium observations on feeding by Melanostigma pammelas (Pisces: Aquarium observations on feeding by Anderson. 1979. and M.E. Belman, B.W. Baird, R.C. and G.Y. Jumper. 1995. Encounter models and deep-sea fishes: numerical simulations and mate Bordes, C., F., R. Wienerroither, F. Uiblein, T.M. Moreno, I.C. Bordes, V.G. Hernández and Moreno, C.C. I.C. Uiblein, Méndez. T.M. Bordes, V.G. 2009. F. Bordes, R. Wienerroither, C., F., Damasceno-Oliveira, A., J. Goncalves, Damasceno-Oliveira, D. Silva, pressuring B. system Fernandez-Duran and A J. Coimbra. 2004. Andriashev, A.P. 1953. A Bagley, A. Bagley, P.M., Smith and I.G. Priede. 1994. Tracking movements of deep demersal fishes in the Porcupine Amorim, M.C.P., Amorim, M.E. M.C.P., Stratoudakis Knight, Y. and 2004. Turner. G.F. Differences in sounds made by courting

Franz Uiblein 17 unea,Y ad . ao. 99 Sm-ira ryhi atvt i de-e bnhc ihs n h By of Bay the in fishes benthic deep-sea in activity rhythmic Semi-diurnal 1979. Ranou. M. and Y.Guennegan, 2003. Stakes. and D.S. fish and deep-sea Goffredi,Schlining egg-brooding S.H. B. of J.C., Aggregations Drazen, Lorance, P., V. Trenkel and F. Uiblein. 2006. Characterizing natural and reaction behaviour of large mid-slope large of behaviour reaction and natural Characterizing 2006. Uiblein. F.P., and V.Lorance, Trenkel Henriques, C., I.G. Priede and P.M. Bagley. 2002. Baited camera observations of deep-sea demersal fishes of the P.M.of and fishes Priede demersal I.G. deep-sea C., of Henriques, observations Bagley.camera Baited 2002. Koslow, J.A., A. Williams and J.R. Paxton. 1997. How many species in the deep sea? A test of a of test sea? A deep the in species fish demersal many How 1997. Paxton. J.R. and Williams J.A., A. Koslow, King, N., P.M. Bagley and I.G. Priede. 2007. Depth zonation and latitudinal distribution of deep-sea scavenging deep-sea of distribution latitudinal and zonation Depth 2007. Priede. I.G. P.M.and N., Bagley King, Isaacs, J.D.andR.A.Schwartzlose.1975. Active animalsofthedeep-seafloor. Scient. American 233:85-91. Lorance, P., S. Souissi and F. Uiblein. 2002. Point, alpha and beta diversity of carnivorous fish along a depth a along fish carnivorous of diversity beta and alpha Point, 2002. Uiblein. F. and Souissi P.,S. Lorance, Koslow, J.A., R. Kloser and C.A. Stanley. 1995. Avoidance of a camera system by a deepwater fish, the orange the fish, deepwater a by system camera a Stanley.1995. Avoidanceof C.A. and Kloser R. Koslow,J.A., diagonal during attitudes horizontal hold Hatchetfishes 1986. Craddock. J.E. and Harbison G.R. J., Janssen, ac, . F Uib F. P., Lorance, Mann, D.A., J. Bowers-Atlmann and R.A. Rountree. 1997. Sounds produced by the striped cusk-eel striped the by produced Sounds 1997. Rountree. R.A. and Bowers-Atlmann J. D.A., Mann, Markle, D.F. and C.A. Wenner. 1979. 1979. Wenner. C.A. and D.F. Markle, Moore, J.A. 2002. Upside-down Upside-down 2002. J.A. Moore, Moku, M., A. Tsuda and K. Kawaguchi. 2003. Spawning season and migration of the myctophid fish Diaphus fish myctophid the of migration and season Spawning 2003. Kawaguchi. K. and TsudaA. M., Moku, Merrett, N.R.andR.L.Haedrich.1997.Deep-demersal fish andfisheries.ChapmanHall. report. progress a animals: mesopelagic of maintenance Aquarium 1976. Anderson. M.E. and J.E. McCosker, Marshall, N.B.1979.Develomentsindeep-seabiology. BlandfordPress,Poole,Dorset. Gigantactinidae). Copeia2002 (4): 1144-1146. theta inthewesternNorthPacific. IchthyologicalResearch50:52-58. M Xenodermichthys copei. Bull. South.Calif. Acad. Sci.75(2):211-219. olsehs atlanticus Hoplostethus 321-331. ih pce: osqecs o cthblt. n Sotn R (d. ep e 20: ofrne n the on Conference 2003: Sea Deep (Ed). R. Shotton, In: catchability. for consequences species: fish Governance andManagementofDeep-seaFisheries.FAO FisheriesProceedings 3(2):162-164 marginatum (Ophidiidae)duringcourtshipand spawning. Copeia1997(3):610-612. migration. Sci.Mar. 68(4):615-619. ontheGordaEscarpment:areproductivehotspot.Biol.Bull.205:1-7. Biscay. Sarsia64:113-116. northeast Atlantic Oceanat15-28°Noff West Africa. Mar. Biol.141:307-314. roughy (Hoplostethusatlanticus descents. J.Mar. Biol. Ass. U.K.66:825-833. method toextrapolatefromlocalglobaldiversity. BiodiversityandConservation6:1523-1532. demersal fishesoftheMid-AtlanticRidge,42to53°N.Mar. Ecol.Prog.Ser. 319:263-274. gradient, Aquat. LivingResour. 15:263-271. nsim a elanostigma n n D Latrou D. and lein tlanticum Copeia1979(2):363-366. Pse: rcihhia) n h By f icy J Mr Bo.As U K 82: K. U. Ass. Biol. Mar. J. Biscay. of Bay the in Trachichthyidae) (Pisces: ih co with ng swimming ). Deep-SeaResearchI42:233-244. . 02 Hab 2002. ite. Evid ns n d on mments ce of of ence r n w a in behavior , be itat, esl spaw demersal ersal spawning in the the in spawning emersal r n clu p colour and haviour pnose hipnose g n h mes the in ning rih (T terns of orange roughy roughy orange of atterns ic zoarcid fish fish zoarcid opelagic : Cer eleostei: alepoceph id fish fish alid Ophidion i: atoidei: 18

Franz Uiblein extreme deep-sea and opelagic fish, mes eep-sea and itions in cond evidence of a light tachowitsch (Eds.), D caused by changes Béland. 1987. Direct ological ellet and P. lein, F., Ott, J. and M. S itats. In Uib Behavioural and morph ater hab H.M. Edenborn, G. Ou ow-w all lander platforms. Oceanogr. Mar. Biol. Annu. Rev. 38:357-392. Annu. Rev. Biol. Mar. lander platforms. Oceanogr. Trans. R. Soc. Lond. B 355: 1299-1303. Trans. im Dunkel. Filander Verlag, Fürth, 181-206. Fürth, 181-206. Verlag, im Dunkel. Filander coast: video, photographic and morphological observations. J. Fish Biol. 69: 970-986. coast: video, photographic and morphological observations. shallow-water habitats: affinities and adaptations. Biosystematics and Ecology Series. 11:91-122. 11:91-122. and Ecology Series. Biosystematics and adaptations. affinities shallow-water habitats: on the design and application of a low-cost hyperbaric chamber. J. Fish Biol. 70: 867-878. on the design and application of a low-cost hyperbaric chamber. sh 195-202. Canada. Env. Biol. Fish. 39: 43-49. Biol. Fish. Canada. Env. Neobythites from the NW Indian Ocean and the Red Sea. Senckenbergiana marit. 23: 109-113. marit. 23: 109-113. Sea. Senckenbergiana Indian Ocean and the Red NW Neobythites from the Melanostigma Melanostigma atlanticum (Zoarcidae) spawning within bottom sediments. Environm. Biol. Fish. 20 (3): Melanostigma Melanostigma atlanticum (Zoarcidae) in muddy bottom sediments of the Laurentian Trough, eastern and bathypelagic scattering layers on the northern Mid-Atlantic Ridge. Deep-Sea Res. 11: 45-58. Ridge. Deep-Sea Res. 11: layers on the northern Mid-Atlantic and bathypelagic scattering ophidioids (Pisces, Brotulidae). Galathea Report 2: 21-48. 2: Galathea Report (Pisces, Brotulidae). ophidioids Ichthyology 39: 110-113. Ichthyology 39: 126-128. tofish: applications of passive acoustics to . Fisheries 31 (9): 433-446. tofish: applications of passive acoustics all, J. 1996. Robison, B.H. 1973. A system for maintaining midwater fishes in captivity. J. Fish. Res. Bd. Can 20 (1): Poulson, T.L. 1971. Biology of cave and deep-sea organisms. National Speleological Society Bulletin 33: 51-61. National Biology of cave and deep-sea organisms. 1971. T.L. Poulson, 74-79. Endeavour 18 (2): of scavenging fishes in the abyss. Tracking Priede, I.G. 1994. Priede, I.G. and P.M. Bagley. 2000. In situ studies on deep-sea demersal fishes using autonomous unmanned Paxton, J.R. 2000. Fish otoliths: do sizes correlate with taxonomic group, habitat and /or luminescence? Phil. Parzefall, Parzefall, J. 2000. Tiefsee-und Höhlenbewohner In: im Vergleich. Uiblein, (Ed.), F, Tiefsee und Höhlen. Leben St. John, J. 2003. Is oyur fish “bent” and will it survive? SPC Live Reef Fish Information Bulletin 11: 31-35. Reef Fish Information Bulletin 11: Live St. John, J. 2003. Is oyur fish “bent” and will it survive? SPC Biol. Fish. 33: 47-52. Food searching decisions in four cyprinid species. Env. 1992. Uiblein, F. Stein, D.L., J.C. Drazen, K.L. Schlining, J.P. Barry and L. Kuhnz. 2006. Snailfishes of the central California Smiley, J.E. Smiley, and M.A. Drawbridge. for 2007. Techniques live capture of deepwater fishes with special emphasis Silverberg, N., Opdal, A.F., O.R. Godø, O.A. Bergstad and Ø. Fiksen. 2008. Distribution, identity, and possible sustaining meso- and Ø. Fiksen. 2008. Distribution, O.R. identity, Godø, O.A. Bergstad A.F., Opdal, Parzef Nielsen, J.G. and Uiblein. Tiefseefische aus 1993. F. Tiefenwasser-und dem new Roten A XVI. species Meer. of Silverberg, Silverberg, N. and L. Bossé. Additional 1994. observations of endobenthic behaviour in the early life history of Robison, B.H. 2004. Deep pelagic biology. J.Exp. Mar. Biol. Ecol. 300: 253-272. J.Exp. Mar. Robison, B.H. 2004. Deep pelagic biology. Rountree, R.A., R.G. Gilmore, C.A. A.D. Goudey, Hawkins, J.J. Luczkovich and D.A. Mann. 2006. Listening Nielsen, J.G. 1977. The deepest living fish, galatheae. A new genus and species of oviparous Sakurai, Y. and K. Kido. 1992. Feeding behaviour of Careproctus rastrinus (Liparidae) in captivity. Jpn. J.

Franz Uiblein 19 ili, . n F Bre. 99 Cmlx rpi itrcin aon oen sad. ca Calne 9 (2): 9 Challenge, Ocean islands. ocean around interactions trophic Complex 1999. Bordes. F. and F. Uiblein, Uiblein, F. 1995. Morphological variability between populations of Neobythites stefanovi (Pisces: Ophidiidae) (Pisces: stefanovi Neobythites of populations between variability Morphological 1995. F. Uiblein, Uiblein, F., F. Bordes, R. Castillo and A. Ramos. 1998. Spatial distribution of shelf-and slope-dwelling fishes slope-dwelling shelf-and of distribution Spatial 1998. Ramos. A. and Castillo R. Bordes, F.F., Uiblein, deep-water demersal in F.,distribution Diversity,Uiblein, depth 1996. and Castillo. F.abundance, R. and Bordes Uiblein, F. 1996a. Anpassungsprozesse bei der Besiedlung lichtreduzierter Lebensräume: verhaltensbiologische, Lebensräume: lichtreduzierter Besiedlung der bei F.1996a. Anpassungsprozesse Uiblein, Uiblein, F. and J.G. Nielsen. 2005. Ocellus variation and possible functions in the genus Neobythites (Teleostei,Neobythites genus the in functions possible and variation Ocellus 2005. Nielsen. J.G. F.and Uiblein, species fish demersal seven of utilization habitat and Behaviour 2003. Latrouite. D. and F.,P.Lorance Uiblein, eel cutthroat northern in behaviour locomotion in Variation 2002. Latrouite. D. and Lorance P. F., Uiblein, ili, . M Yuglt, . aoy F Pgs M Pcea ad . osy 20. n iu bevtos of observations situ In 2006. Gorsky. G. and Picheral M. Pagès, F. Jacoby, C. Youngbluth, M. F., Uiblein, Uiblein, F. 1996b. Behavioural and morphological plasticity in facultative cave dwellers and secondary deep-sea secondary and dwellers cave facultative in plasticity morphological F.and Uiblein, Behavioural 1996b. Uiblein, F. 1996c. Constraints and exploratory windows in light-reduced . In: Uiblein, F., J. F., Uiblein, In: habitats. marine light-reduced in windows exploratory and Constraints 1996c. F. Uiblein, Uiblein, F. (Ed.).2000. Tiefsee undHöhlen.Lebenim Dunkel. Filander, Fürth. Uiblein, F., J.P. Durand, C. Juberthie and J. Parzefall. 1992. Predation in caves: the effects of differential prey differential of effects the caves: in Predation 1992. Parzefall. J. and Juberthie C. J.P.F.,Durand, Uiblein, Uiblein, F., W. Klausewitz and J.G. Nielsen. 1994. Depth dependent morphological variation in two ophidiiform two in variation morphological dependent Depth F.,1994. Uiblein, Nielsen. J.G. W. and Klausewitz mechanical by prey mobile detect anguinus Proteus salamander cave the Does 1993. Parzefall. F.J. Uiblein, and Uiblein, F.,Uiblein, 1995. Parzefall. J. Engelkeand S. Trade-offbetween patch detectability andnutrientcontentin visual collected bybottomlonglineoff LanzaroteandFuerteventura, CanaryIslands.Mar. Ecol.19:53-66. choice ofthePyreneansalamanderEuproctusasper. Ethology, 101:39-45. fishes off LanzaroteandFuerteventura,Canary Islands. J.FishBiol.49A:75-90. from thedeepRedSeaandGulfof Aden. Mar. Ecol.Prog.Ser., 124:23-29. on theBayofBiscaycontinentalslope,NE Atlantic. Mar. Ecol.Prog.Ser. 257:223-232. (Synaphobranchus kaupi)ontheBayofBiscaycontinental slope.Deep-SeaResearchI49:1689-1703. 15-16. klgsh ud mor und ökologische deep-water fishes in four canyons off the Georges Bank, NW Atlantic. In: Shotton, R. (Ed). Deep Sea Deep (Ed). R. Shotton, In: Atlantic. NW Bank, Georges the off canyons four in fishes deep-water Ophidiidae). IchthyologicalResearch,52(4):364-372. Tiefseefischen. Habilitationthesisintenpublications,UniversityofSalzburg. 03 Cneec o te Go the on Conference 2003: Proceedings 3(1):98-106. n M Sahwtc (d.: epsa n etee hlo-ae hbtt: fiiis n adaptations. and affinities habitats: shallow-water extreme and Deep-sea (Eds.): Stachowitsch M. and fishes: adaptivestepsintothedark.Mém.Biospéol.,22:211-214. Biosystematics andEcologySeries, 11: 165-182. oiiy n drns o te oaig eaiu o to aaadr, urcu apr n Proteus and asper Euproctus salamanders, two of behaviour foraging the on darkness and mobility cues? Mém.Biospéol.,20:261-264. anguinus. Behav. Process.,28:33-40. fishes from the deep Red Sea: evidence for species-specific structure in vertical distribution. Cybium 18: Cybium distribution. vertical in structure species-specific for evidence Sea: Red deep the from fishes 15-23. lgsh U phologische nne n Man and vernance nte cugn n hö an rsuchungen t f D of agement hlen a Fi eep-sea bewoh e S nenden sher . A Fi FAO ies. andern und und alamandern eries sheries

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